Document 12787087

advertisement

Tech. Report 14, 155 p. (1962). (7) S..,EoEcoR, G. W.: Statistical methods. 5th Edition. Ames, Iowa, Iowa State University Press.

534 p. (1956). (8) SQUILLACE, A. E., EcHoLs, R. M., and DoRMAN, K.

M.: Heritability of specific gravity and summerwood per cent and relation to other factors in slash pine. Tappi 45: 599-601 (1962).

(9) VAN BuiJTENEN, J. P., JoRANSON, P. N., and MAcLAURIN, D. J.:

Pulping southern pine increment cores by means of a small-scale kraft procedure. Tappi 44: 166-169 (1961). (10) vAN BUIJTENEN, J. P.,

ZoBEL, B. J., and JonANsoN, P. N.: Variation of some wood and pulp properties in an even-age loblolly pine stand. Tappi 44: 141-144

(1961). (11) vAN BuiJTENEN, J. P.: Heritability estimates of wood density in loblolly pine. Tappi 45: 602-605 (1962). (12) WINK, W.

A., and VAN EPEREN, R. H.: The development of an improved zero­ span tensile test. Tappi 45: 10-24 (1962). (13) ZoBEL, B., THoR­

BJORNSEN, E., and HENSON, F.: Geographic site and individual tree variation in wood properties of loblolly pine. Silvae Gene~ica 9:

149-176 (1960). (14) ZoBEL, BnucE, CoLE, DoNALD, and SToNr:CYPIIER,

RoY.: Wood properties of clones of slash pine. Pro c. Forest Gen­ etics Workshop, p. 32-39., SAF, Macon, Georgia, October 26,

1962.

Inheritance of Needle and Bud Characteristics of Slash Pine

By FRANK c.

SORENSEN

(Received for publication December 3, 1963)

The purpose of this investigation was to analyze the ex­ tent of gene control of several variable needle and bud characteristics in slash pine (Pinus elliottii ENGELM.). Var­ iation in needle and bud characteristics, particularly needle length, has often been used as a tool in investigations of geographic variation and of interspecific hybridization in pine species. Only rarely, however, have the observations and studies included determination of an approximate de­ gree of genetic control of variation, or compared different characteristics on this basis.

Review of Literature

DENGLER (1938), in studying a number of pines, found that different races still varied in needle length in the second generation. In another European investigation, BuRGER

(1941) found that Scots pines (P. sylvestris L.) of different origins, but grown on the same site, required different amounts of needles to produce the same wood volume in­ crement.

Three pertinent observations have been made in ponde­ rosa pine (P. ponderosa LAws.) in this country. LoRENZ

(1949), in a seed source study, reported major differences in color and texture of needles of trees of different sources when grown on the same site. In a similar study, Mu~GER

(1947) noted that progeny from some seed sources were bushier than others and that variation existed in needle color, duration of needle retention, and angle of the needle with the twig. Average needle length ranged from 4 inches for one source to 7 !~ inches for another. WEIDMAN (1939) showed that 22- to 26-year-old ponderosa pine progenies from different geographic sources differed in the number of needles per fascicle, length of needles, and general ap­ pearance of foliage. These differences corresponded to dif­ ferences between trees of the various parent localities, and the conclusion was drawn that the characteristics were strongly inherited.

Several needle characteristics of slash pine and some of its hybrids were investigated by MERGEN (1958). The num­ ber of resin ducts per needle and the number of stomata

1

)

Based upon a thesis presented by the author in partial fulfill­ ment of the degree of Master of Science in Forestry at the Uni­ versity of Florida, 1960. The investigation was conducted in co­ operation with the Southeastern Forest Experiment Station, Lake

City, Florida, which supplied the plant material. Acknowledge­ ments are due to Dr. T. 0. PERRY for suggesting the problem and to Mr. A. E. SQuiLLACE for much helpful advice. Further acknow­ ledgements are made to Drs. R. R. SILEN and H. IRGENs-MoLLER for reviewing the paper. The author is presently a Research Forester with the Pacific Northwest Forest & Range Experiment Station at

Corvallis, Oregon, USA. per millimeter of length showed a pattern which could be correlated with the geographic distribution of the species.

The number of teeth on the needle margin, however, did not appear to be so correlated.

S....>.

01=::

Q)Q)

1=::0.0

..........

UA

G-1

G-2

G-3

G-4

G-7

....

Q)s....>.

Ace

UlQ)Q)

Q)~=::O.O

2::£8

~<:.>A

Q)O.O

Q)i=:: z~

Number mm.

1X2 14

1 X 7 14

3X2 14

4 X 1 14

2 X 7 2

)

7

3 X 4 2

)

4 X 2 2

)

7

4

6

8

2

6

1

263

239

248

264

297

244

248

227

252

252

-1)

240

Q) ......

Procedure

Description of Material

The plant materials used in this study were: 1) 15-year­ old rooted cuttings obtained from a group of slash pine trees selected for their high gum-yielding ability, and 2)

15-year-old progenies produced by controlled pollinations among the same high-gum-yielding selections. Both of the plantations were situated near Olustee, Florida. The ma­ terial was made available to the author by the Lake City

Research Center of the U.S. Forest Service.

The clones were growing in an informal plantation about

3/4 acre in size, containing varying numbers of ramets from each clone. The ramets of some clones were planted in un­ replicated row plots while those of other c1ones were located more or less at random. Five of the clones were chosen for study (Table 1).

The progeny plantation was about 6 acres in size. In it, varying numbers of trees of most progenies were planted randomly within seven blocks. A few progenies were re­ presented only in replacement blocks where the trees were planted in unreplicated row plots. Seven of the progeny

Tabte 1. Clonal and progeny a•;erages for all traits measured. oQ)Q)

E

:tj'gE

Q)::lo

Z.c:>

Ul

Q,)

:tj

~

'0

Q)s....l=::

Q)Q);:l

Z

A.C

Q)

~.s::.s::

•I"'"!

;-I~

<:JCUo.o

UlQ)r;:

<U.s::

Q)

I":<.Ul ......

2·(1)

~~~ z~~

1

)

2)

Not measured.

Progenies represented in the replacement blocks only.

Q)

Cil

~:S

'00.0

;::li=::

~~ cc. Number mm. mm. mm.

CLONES

.785

.520

.667

.632

.786

2.17

2.04

2.00

2.05

2.30

14.1

105

13.5

16.0

17.0

11.4

15.2

5.7

17.8

9.9

14.1

11.5

13.1

15.2

-1)

PROGENIES

- - - - - · - ­

- - - ­

.515

.509

.472

.557

.505

-1)

.454

2.13

2.14

2.04

2.31

2.12

2.14

2.05

11.6

13.3

11.2

13.0

-1)

-1)

12.3

----~

16.3

14.9

12.6

19.6

15.8

16 9

17.3

-1)

-1)

14.0

15.7

-1)

15.9

-1)

109

groups were chosen for study. Of these, four were within the replicated blocks while three were within replacement blocks (Table 1).

Foliage and bud collections were made in late January and early ,February, 1959. At this time the terminal vege­ tative buds of the upper 2 or 3 whorls of primary branches were about 3 to 5 centimeters long. Visible elongation of the new needles had not yet begun. A total of 10 needle bundles were taken from each tree. Five of these were selected from branches in the middle one-third of the crown and five from branches in the lower one-third. The selection of bundles was restricted to 1- and 2-year-ol.d wood only. Vegetative bud samples were taken from the extreme upper portions of the crown. Two buds of vigorous appearance were taken randomly from the top two whorls of each tree.

After collection the needles and buds were placed in polyethylene bags containing a small amount of water and stored in a refrigerator at about 40° F. until measurement.

Measurements were then begun and completed within about 6 weeks. The traits measured are listed below.

Needle length was measured from the base of the needle bundle to the tip of the needles.

Needle-bundle volume was measured by water displace­ ment and expressed in relative terms. The bundles were immersed in a narrow glass tube containing water and the length of displacement of the column of wate:· was me:~s­ ured. All needles were measured within 2 weeks after storage. Through a "uniformity trial" the effect of length of storage time upon clonal or progeny differences was shown to be not greater than 3 percent.

Fascicle sheath length was measured from the base of the needle bundle to the tip of the sheath.

Needle "divergence" as used here, refers to the separa­ tion of needles as they extend out from a single fascicle sheath. The distance between needles of the same bundle was measured in millimeters at a point 10 centimeters from the base of the fascicle sheath.

Bud scale length was measured on scales located ap­ proximately 1 centimeter back from the tip of the bud. Two scales were plucked from opposite sides of the bud at this point.

Statistical Analysis

Because the materials were growing in two plantations and represented two different types of propagation, the presence of genetic effect could be tested through: 1) vari­ ance of individuals and clones in the clonal plantation, 2) variance of progenies in the progeny plantation. 3) corre­ lation between parents {represented by clones) and pro­ genies.

Variance of the clones was analyzed according to a hier­ archal classification model

(SNEDECOR,

1956). The main samples were clones. The clo!les were composed of sub­ samples of ramets which in turn were composed of sub­ samples of needle bundles. These classifications were used for all traits except needles per bundle, which could only be classified into clones and ramets. The sources of Vlria­ tion and the estimated variance parameters were:

Source of variation Parameters estimated

Clones

Trees in clones a2

+ naT 2 + bnac 2 a2

+ naT2

Bundles in trees in clones a2 where: a2 is the component of variance due to bundles of the same tree, oT2 is the component of variance due to trees of the same clone, oc 2 is the component of variance due to clones, n is the number of bundles (buds) on which the indi­ vidual tree average is based (10 for needle traits and

2 for the bud traits), b is the number of trees on which the clonal mean is

'based (4.22 for needle characteristics, and 5.21 for the bud trait). :Because the clones contain different num­ bers of ramets, the averages were computed using the following formula for unequal sample sizes where:

(SNEDECOR,

1956): b = _1a - 1

(c -l:bi2) c a is the number of clones, c is the total number of trees in all clones, bi is the number of ramets in each clone.

The components of variance for clones and trees within clones were then determined.

Because trees of the different clones have different geno­ types, it can be said that there is genetic control of the variation of a particular trait if the component of variance contributed by clones is significant.

Data from the randomized block design of the progeny plantation were subjected to the typical analysis of vari­ ance. The sources of variation and the estimated para­ meters were:

Sources of variation

Blocks

Crosses

Remainder

Parameters estimated a2 a2

+ akB2

+ ckc2 a2 where: a 2 is the component of variance due to trees of the same cross or parentage, kc

2 is the component of variance due to crosses, kn 2 is the component of variance due to blocks, a is the number of crosses on which the block estimate is based (4 for needle traits and 2 for bud traits), c is the number of blocks on which the cross estim~te is based (7).

If genetic influence is measurable, trees cf the s::1me parentage on the average should resemble each other more than they resemble individuals of other parentages. The evidence in the analysis for this is a significant component of variance due to crosses.

Parent-progeny correlations were determined between the clones and their progenies. The progeny value for each comparison was estimated by the average value of all trees of the progeny. The parental value for each comparison was estimated by the mid-point value of the p2rents mak­ ing up the cross. Each parental value was estimated by the average of the ramets which represented it ir; the clonal plantation. In terms of this analysis then, genetic effect is present if the parents and their progeny show positive correlation. Regressions of the offspring on the mid-point of the parents were also determined.

110

~. m ~ ~ fr.

~C;~'Ot:!:~

>--"• (!)

~

~Q)o-o.::s~~

2ro~~o.'[

::SO.~O.H.(i) s+

0 roro;=;Cil

'"1

(ll

0

,;::g .... roe:."-+>

.,.... Q)

~

::r-

...... §

'"1

~ ~

> - ' ( )

O"Q) .,..

Q)

0

CX) ,;:: I-"•

~ c:so[fii~

~'Oro(D-(i)

...... 0

(!)

~ ~

'<::::S>-1.,....~-S,

(!) () sg.~ro(;~

~ t-i>

(!)

::r-

......

0" c::

-~ ~ ~

,_. 0 0

::s . (i)

~

8'"1H.~I-lJ·~ !:ll s

~ 8" r.;· ;:; t-3 roroor~ ::r­

Q)O.....-oe?..ro

'"1 ,_.

(!) (!)

'"1 ......

.,.... ()

0.(/l~S.g.Q' e.

.,....roro§Q)::l

0

::r-

(!)

~-:::~ro~m

0'"

,;::

::s

'"1

>--"•

§ f)

0

::s

::;;

~.

1.1

(10.

>--"•

::s

~

(ll ro ro o: r.;· r.;·

~~t-3::trog.

(!)

I

Q)

::r-

00 (!)

,;::

0

(ll

~

Q)

......

::l

I

'<:

Tabte 2. Analyses of variance for all traits measured in the clonal plantation

Characteristic

Needle length

Bundle volume

Needle divergence

Fascicle sheath length

Needles per bundle

Bud scale length

Degress of freedom

Clones Trees Needles

4

4

4

4

4

3

18

18

18

18

18

18

207

207

207

207

_2)

22

Clones

Mean squares

Trees

11,950**

4,407**

722.0**

307.8**

.0300*

36 10**

517.6*

67.80

27.82

2.528

.00833

1.415

Needles

291.0

74.64

19.74

1.652

1.732

I

Estimated variance components

Clones Trees Needles

270.9

102.8

16.45

7.234

.00691

3.329

22.66

Negative

1

)

.808

.0866

.00833

Negative

1

)

291.0

74.64

19.74

1.652

1.732

I Clone contribution3)

Percent

84

94

86

97

45

82

1) If the variance component was estimated as a negative value, it was treated as zero for determining the estimated clone contribution.

2) Estimate of individual needles was not possible. nac2

3) Clone contributions obtained by the formula, in which "n" is the number of needles on which the tree average is based, and the variances ae2

+ nat2

+ nac2 are those due to error, trees, and clones as shown by the subscripts.

** Significant at the .01 level of confidence.

* Significant at the .05 level of confidence.

~

~

~

J8"~H

~

0

::s

;:::;: ~ '0 .,....

~

<

p;· ~ ~

::::::.: ::s Q)

(ll

;:::;: ()

!"

(ll

Q)

~s

0

(!)

~

Q)

,;::

.,....

I-"• s

::s

(!) s

'<:(J)Q)Q) . ~ !"

::

(!)

§'

Q)

: ; (!)

~

(!)

(!) ()

Q)

Q)

(ll

<

(ll

(!)

~'"1

.,....

(ll I-"•

::s

::rH.

0 .,....

0"~

(!)

()

Q)

(!)

'"1

::r­

0

(10.

(ll

(!) (!)

,;:: ::s ::s

0

(ll

(!)

~

(!)

'"1

(!)

~

::r-

(ll

~

.,.... (ll

::r-m n"~

............ g

(!)

::s

Q)

......

'<

(!)

Q)

(jJ

~

0

~ ro

,.,.. (10.

•v .,.... C""+

()?"

(ll g

(!)

(ll r.;· g.

~ o.:;..

I-"•

(10. ,_.

::s '<

I-"• (ll

H. .....

~'<

(ll

::s

.,....

'"1

(ll

Q) 0

H.

0.0

::+;g.

'"1 f1) 0

::s ::s

~

(!)

( ) o-::t

(!)

Q)

C""+'-1 g ~H. ~ Q)

(!) () s r.;·.,....

(!) .,....

:;~()~

<? ~ 0

(10.

'< ::s

(ll

.,.... ,;::

~ ~

'<

~.

g'

0 (!)

0"

(!)

.,....o,,;::,o

'OH>Oro

(!)

(ll

0

::s ::s

'"1 (ll .,....

::s

.,.... (ll

S.

'"1 ­

.....

~()

(ll

::s

.,.... (!)

::r-2::

(!) (!)

Characteristic

Needle length

Bundle volume

Needle divergence

Fascicle sheath length

Needles per bundle

Bud scale length

Tabte 3. Analyses of variance for all traits measured in the progeny plantation

1------------------­

Estimated variance components

Degrees of freedom

Crosses Blocks Remainder Crosses Blocks Remainder I Crosses Blocks Remainder

3

3

3

3

3

1

6

6

6

6

6

6

18

18

18

18

18

6

855.5**

56.12*

57.96*'

7.699**

.0446

10.81*

127.1

13.42 g,208

.3155

.0569

1.738

75. )4

12.40

8.608

.4777

.0240

1.408

I

I

I

114.4

6.25

7.05

1.03

.00294

1.34

1

)

Cross contribution obtained by dividing the

** Significant at the .01 level of confidence.

* Significant at the .05 level of confidence. cross cumponent by the sum of the cross plus remainder components.

13.86

.255

Negative

Negative

.0082:2

.166

75.64

12.40

8.608

.4777

.0240

1.408

I

Cross contribution!)

Percent

60

34

45

68

11

49

260 NEEDLE LENGTH

250

240

230

220 r

2

=.584 b

= .540

240 250 260 270 280

5C

BUNDLE VOLUME

35 r

2 = .465 b

= .324

45 50 55 60

(tHlliJ:,eters of" lic:tuLl d.L'[J.Cacer·,ent in t_£.L::L.;:...; ;:,eu:-=.uring tuUe)

65

20 NEEDLE DIVERGENCE

18

16

14

12

10 12 14 r

2

= .555 b = .748

16 18

2.4 NEEDLES PER BUNDLE

2.3

2.2

2.1 o3•4

~

_..----'ix2 °2x7

0

4x2

0

3x2 r

2

=.097

b =.334

2.0

2.0 2.1 2.2 2.3

(Average nwnber of needle&•·!<er bw!d Le)

2.4

FASCICLE SHEATH LENGTH

17 BUD SCALE LENGTH

14

16

13

15

12

14

/

0

3x2

II b

=

.549 13 r2 =.916 b =.806

12 13 14

(Millimeters)

15 16 12 13 14

(tUllimeters)

15 16

Figre 1. Scatter diagrams and regressions of pro•geny on parents for the various traits. The ordinates are scaled for the progeny values; the abscissae are scaled for the parental values.

See text for further description of traits and measurement techniques.

were not randomly selected, the term "heritability" has ranged from 34 to 68 percent of the cross plus remainder not been used. components.

Analyses of variance of progeny:

The crosses varied sig­

Parent-progeny correlation analyses:

Because few de­ nificantly for all measurement traits but not for needles grees of freedom were available, an extremely high cor­ per bundle

(Table 3).

Because the variance of trees within relation was necessary to achieve statistical significance. a cross is greater than the variance of ramets within a Only one trait, fascicle sheath length, attained this signifi­ clone, the genetic contributions were lower than in the cance. The squares of the correlation coefficents showed clonal analyses. Except for needles per bundle they still the estimated additive parental contributions to the proge­

112

ny performances. They ranged from 10 percent for needles per bundle to 97 percent for fascicle sheath length (Table 4 and Figure 1).

Table 4. Parent-progeny correlation coefficients and regression coefficients for various traits.

Characteristic

Needle length

Bundle volume

Needle divergence

Fascicle sheath length

Needles per bundle

Bud scale length

** Significant at the .01 level of confidence.

Correlations between traits: It is often of value to know, particularly in selection work, if traits are negatively or positively correlated. Although the characteristics examin­ ed in the present study are not economically important, their relationships were determined using the material in the clonal plantation (Table 5). The following formula was applied to correct for unequal numbers of individuals within the clones 2). r=

I

I

Degrees of freedom

4

I

I

4

5

3

5

1 r2 r b

0.584 0.764 0.540**

.465 .682 .324**

.555 .745 .748**

.970 .985** .549**

.097 .312 .334

.916 .957 .806**

Lni·(xi-x)(Yi-Y)

{L 0 i (xi- x )

2

I

{L 0 i (Yi- Y)

2

I

Table 5. Between trait correlation coefficients

Needle length

Needles per bundle

Bundle volume

Fascicle sheath length

.696 .777

.792

* Significant at the .05 level of confidence.

** Significant at the .01 level of confidence.

Fascicle [ Bud sheath \ length scale length

.904*

.405

.665

.968*

.349

.658

.995**

Discussion

All traits except needles per bundle behaved in a man­ ner consistent with a high degree of genetic control. Based on the results of all analyses, fascicle sheath length seems to be the most strongly inherited, followed by bud scale length.

Needle length is subject to considerable variation within the genotype as shown by the significant within clone dif­ ference and by the high estimated component of variance due to needles of the same tree (Table 2). The high variance of the latter possibly explains the former. Probably this trait is somewhat more susceptible to environmental in­ fluences such as position on the branch, position on the tree, age of needle, etc., than are the other traits. If this is true, and small samples (10 bundles) are taken, the effect of the within tree influence could easily be estimated differently in one ramet than in another. A larger sample, or greater restrictions on sampling, should decrease the within tree and remainder variance terms.

The same reasoning applies to bundle volume, at least to the extent that volume is dependent upon length. Although this trait shows no significant difference between trees of the same clone, the component of variance due to needles is fairly high (Table 2). Also, the remainder component for crosses (Table 3) is higher than that of any other measure­ ment trait.

Needle "divergence" has a high genetic component of variance, but it also has one complicating feature involved

2) Personal communication from Mr. K. W. DoRMAN. in its measurement and inheritance. The extent of "diver­ gence" apparently depends primarily on relative growth rate of certain parts of the needle, and the relative growth rate of these parts can be modified by other needle struc­ ture characteristics. For example, one clone (No. 2) had twisted needles and all the progeny which had that clone as a parent ranked lower on the "divergence" scale (see

Figure 1) than was predicted. (The predicted value was based on the trend established by progeny which did not have "twisted" as a parent.) Therefore, some clones and progeny groups are remarkably uniform in needle "diver­ gence", but in general, considerable between tree variation exists, particularly in the progeny plantation.

Needles per bundle has a significant, but relatively small, genetic component in the clonal test, and a nonsignificant genetic effect in the other two analyses. The reason for this is the extreme variability among individuals in some of the clones. That is, the ramets of one clone are exclusively two­ needled; the ramets of another clone vary from entirely two-needled to 40 percent three-needled. In the progeny plantation the variability is even more striking. The 14 offspring in one progeny group are made up of 5 trees having 90 percent 2-needle bundles and 9 trees having 100 percent 2-needle bundles. In contrast, the 14 offspring of another mating include one tree which is 100 percent 3­ needled, 3 trees which are 60 to 80 percent 3-needled, 7 trees 10 to 30 percent 3-needled, and 3 trees 100 percent

2-needled. Thus, it appears that some gene combinations exert strong control over number of needles per bundle, but that others are less effective.

Generally, the parent-progeny correlation and regression results are quite compatible with results from the analyses of variance. The main exception is bud scale length. It ap­ pears to be under considerably more genetic control when parents and progeny are compared (Table 4) than when the progeny alone are analyzed (Table 3).

In conclusion, the limits of variation of all the traits except needles per bundle are under strong genetic control.

This conclusion is based on r(1) the cross contribution to the variation in the progeny plantation and (2) the cor­ relation and regression coefficients from the parent-progeny correlations. Fascicle sheath length and bud scale length are found to be the most strongly inherited, needle length and needle divergence are second, and bundle volume third.

Summary

Fifteen-year-old slash pine, growing in a clonal and in a progeny plantation were investigated to determine if, and to what extent, several characters of needles and buds were inherited. The characters were needle length, needle bundle volume, fascicle sheath length, needle "divergence", num­ ber of needles per bundle, and bud scale length. Three sta­ tistical analyses were applied: the variances of the clones and ramets were analyzed, the variances of the progenies were analyzed, and parent-progeny correlations were de­ termined. All traits except number of needles per bundle were found to be strongly controlled by the genotype; the genetic control of the latter, although significant in one analysis, was relatively slight.

Resume

Titre de l'article:

Heredite

des caracteres d'aiguiLles et de bourgeons ·chez Pinus elliottii.

On a etudie des Pinus elliottii de 15 ans, poussant dans un pare a clones et dans un test de descendance, afin de

113

determiner si et dans quelle mesure plusieurs caracteres d'aiguilles et de bourgeons sont hereditaires. Ces caracteres etaient: la longueur des aiguilles, le volume des fascicules d'aiguilles, la longueur de la gaine, la «divergence» des ai­ guilles, le nombre d'aiguilles par fascicule, et la longueur de l'ecaille du bourgeon. Les trois analyses statistiques suivantes ont ete effectuees: variance des clones et des ramets, variance des descendances, et correlations parent­ descendant. Taus ces caracteres, excepte le nombre d'ai­ guilles par fascicule, se sont rev~eles fortement contr6les par le genotype; le contr6le g~€metique du dernier caractere, bien que significatif dans une des analyses, est relative­ ment faible.

Nadel-,Divergenz", die Anzahl der Nadeln je Biiridel und die Lange der Knospenschuppen. Drei statistische Analysen wurden angewandt: Die Varianzen der Klone und Klon­ glieder wurden analysiert, ebenso die Varianzen der Nach­ kommenschaften und die Eltern-N a chkommenschafts-Kor­ relationen bestimmt. Alle Merkmale, ausgenommen die Na­ delzahl je Biindel, wurden durch den Genotyp stark kon­ trolliert; die genetische Verankerung der Nadelzahl je Bun­ del war mit einer Ausnahme nur relativ schwach.

Zusammenfassung

Titel der Arbeit: Vber die Vererbung von Nadel- und

Knospenmerkmalen bei Pinus elliottii.

15jahrige Kiefern aus Klonplantagen und Nachkommen­ schaftsanbauten wurden untersucht, urn festzustellen, ob und in welchem Umfang einige Nadel- und Knospenmerk­ male vererbt werden. Untersucht wurden die Nadellange,

Grol3e des Nadelbiindels, Lange der Faszikelscheide, die

Literature Cited

BuRGER, H.: Holz, Blattmenge und Zuwachs. v. Fichten und Foh­ ren verschiedener Herkunft auf verschiedenen Kulturorten. Mit­ teil. Schweiz. Anst. Forstl. Versuchsw. 22: 10-62 (1941) DENGLER,

A.: Fremde Kiefernherklinfte in zweiter Generation. Zeitschr.

Forst- u. Jagdw. 70: 150-162 (1938).- LoRENz, R. W.: Ponderosa pine seed source study. Ill. Agr. Expt. Sta. Forestry Note 4, 3 pp. (1949).

MERGEN, F.: Genetic variation ia needle characeristics of slash pine and some of its hybrids. Silvae Genetica 7: 1-4 (1958). -

MuNGER, T. T.: Growth of ten regional races of ponderosa pine in six plantations. Pacific NW. Forest & Range Expt. Sta. Res. Note

39, 4 pp. (1947). SNEDEcoR, G.

w.:

Statistical methods. Iowa State

College Press, Ames, Iowa. 534 pp. (195'6). WEIDMAN, R. H.:

Evidences of racial influence in a 25-year-old test of ponderosa pine. Jour. Agr. Res. 49: 855-887 (1939).

Inheritance of Yellow Oleoresin and Virescent Foliage in Slash Pine

By JoHN F. KRAUS and A. E. SQUILLACE 1

)

(Received for publication November 26, 1963)

Simply inherited, qualitative traits, although often not of economic importance, can be valuable aids in studies of reproductive systems, cytogenetical analyses, population studies, and others. Such "gene markers" hav2 been used extensively in animal and crop plant genetics, and results of these studies have been basic to breeding work. In for­ est trees, where life spans are relatively long, qualitative traits are especially useful when their expression occurs early in the life of the plant.

The occurrence of two such characters in slash pine (Pinus elliottii ENGELM.) are described in this report. One of these is yellow oleoresin, and the other is a type of chlorophyll deficiency, "virescent foliage". They were found incidental­ ly in a program designed to breed for high oleoresin yield

(MITCHELL et al., 1942). A third qualitative character oc­ curring in slash pine, albinism, was reported upon earlier

(SQUILLACE and KRAUS, 1963).

Crosses were made in the spring of 1960 (table 1). At the time of cone collection, cones pollinated by windborne pol­ len also were collected. Seed were extracted, and wings removed. Sound and unsound seed were separated in an air column seed separator. Seed were sown in the spring of 1962 in row plots randomized and replicated three times.

In November 1962 a narrow strip of ba1:1k and wood ap­ proximately 1 inch long was cut from the stem of each seedling. Within 2 hours enough oleoresin had exuded from the wounded surface to permit discrimination of oleoresin color.

Table 1. Results of crosses on slash pine 29 producing yellow oleoresin.

Mating

29 X 29

29 X 1960 mix

29 X wind

1

)

I

Seedlings producing Total seedlings yellow oleoresin examined

15

0

1

Number

15

271

140

1) The "1960 mix" consisted of equal volumes of pollen from

21 trees.

Yellow Oleoresin

Normal slash pines yield an oleoresin that is transparent and colorless when fresh. Occasionally deviants from this type have been found. One, identified as tree 29, yields a golden yellow oleoresin. Analysis of tl-J.e turpentine frac­ tion of this oleoresin by gas chromatography has revealed no difference in chemical composition which would account for the unusual color. However, the {3-phellandrene con­ tent was unusually high, 40.5 percent in contrast to 7 per­ cent or less in commercial turpentine reported by FISHER et al. (1957). Turpentine yield as a percent of the oleoresin weight was relatively low, 15 percent, compared with the normal yield of about 22 percent given by MIRov (1961) 2

).

') Research Foresters, stationed at the Naval Stores and Timber

Production Laboratory, Southeastern Forest Experiment Station,

Forest Service, U. S. Department of Agriculture, Olustee, Florida.

All of the selfed seedlings produced yellow oleoresin.

All of those from outcrosses produced normal colorless oleoresin (table 1). One seedling among the wind-pollinated progeny produced yellow oleoresin. This probably result­ ed from natural self-pollination.

These results indicate that tree 29 is homozygous for a recessive gene or genes governing production of yellow oleoresin. 1Further breeding by selfing or crossing trees

2) Thanks are due Mr. GoRDON

s.

Fisher of the U. S. D. A. Agri­ cultural Research Service, Naval Stores Laboratory, Olustee,

Florida, for his analysis and interpretation of the oleoresin com­ position.

114

Download